21
Paleobathymetry in the backstripping procedure: Correction for oxygenation effects on depth estimates D.J.J. van Hinsbergen a, T , T.J. Kouwenhoven b , G.J. van der Zwaan b a Vening Meinesz Research School of Geodynamics (VMSG), Utrecht University, Faculty of Geosciences, Budapestlaan 4, 3584 CD Utrecht, The Netherlands b Institute for Paleoenvironments and Paleoclimate Utrecht (IPPU), Utrecht University, Faculty of Geosciences, Budapestlaan 4, 3584 CD Utrecht, The Netherlands Received 25 February 2004; accepted 23 February 2005 Abstract This paper aims to provide a straightforward and easily applicable method for estimating the depositional depth evolution of marine basins. Vertical movements of the basin floor can be reconstructed from the sedimentary record, and more accurately constrained when information from the sedimentary history is combined with palaeodepth estimates derived from fauna. To this end we propose to extend an existing method based on the percentage of planktonic foraminifera with respect to the total (planktonic and benthic) foraminiferal association, which is expressed as the percentage planktonics (%P). The ratio between planktonic and benthic foraminifera is related to water depth, and the %P generally increases with increasing distance to shore. However, next to water depth the oxygen level of bottom waters has a profound effect on the abundance of benthic foraminifera, and as such influences the %P. Depending on basin configuration, the oxygen level at the sea floor can vary on Milankovitch time scales and is reflected by the fraction of benthic foraminiferal species that indicate an effect of oxygen stress on the biotic system. These species can be used as stress-markers and their percentage with respect to the total benthic population is here expressed as %S . To assess whether the effect of sea-floor oxygenation impairs depth reconstructions, we studied the percentage of planktonic foraminifera (%P) in five well-dated sedimentary successions from the Lower Pliocene of Crete, Corfu and Milos in Greece. Additionally, we assessed whether different foraminiferal size fractions and counting methods affect the determination of the percentage of planktonic foraminifera. The palaeobathymetric evolution calculated for each basin was confirmed for all successions by an independent check on depth-related occurrences of benthic foraminifera. After correction for bathymetry changes of the basin due to sedimentation, compaction and eustatic sea level variations, the 0031-0182/$ - see front matter D 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2005.02.013 T Corresponding author. Present address: Geology Department, University of Leicester, University Road, Leicester LE1 7RH, UK. Tel.: +44 116 2523629; fax: +44 116 2523918. E-mail address: [email protected] (D.J.J. van Hinsbergen). Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245– 265 www.elsevier.com/locate/palaeo

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Page 1: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

www.elsevier.com/locate/palaeo

Palaeogeography, Palaeoclimatology, Pa

Paleobathymetry in the backstripping procedure: Correction for

oxygenation effects on depth estimates

D.J.J. van Hinsbergena,T, T.J. Kouwenhovenb, G.J. van der Zwaanb

aVening Meinesz Research School of Geodynamics (VMSG), Utrecht University, Faculty of Geosciences,

Budapestlaan 4, 3584 CD Utrecht, The NetherlandsbInstitute for Paleoenvironments and Paleoclimate Utrecht (IPPU), Utrecht University, Faculty of Geosciences,

Budapestlaan 4, 3584 CD Utrecht, The Netherlands

Received 25 February 2004; accepted 23 February 2005

Abstract

This paper aims to provide a straightforward and easily applicable method for estimating the depositional depth

evolution of marine basins. Vertical movements of the basin floor can be reconstructed from the sedimentary record, and

more accurately constrained when information from the sedimentary history is combined with palaeodepth estimates

derived from fauna. To this end we propose to extend an existing method based on the percentage of planktonic

foraminifera with respect to the total (planktonic and benthic) foraminiferal association, which is expressed as the

percentage planktonics (%P).

The ratio between planktonic and benthic foraminifera is related to water depth, and the %P generally increases with

increasing distance to shore. However, next to water depth the oxygen level of bottom waters has a profound effect on

the abundance of benthic foraminifera, and as such influences the %P. Depending on basin configuration, the oxygen

level at the sea floor can vary on Milankovitch time scales and is reflected by the fraction of benthic foraminiferal species

that indicate an effect of oxygen stress on the biotic system. These species can be used as stress-markers and their

percentage with respect to the total benthic population is here expressed as %S.

To assess whether the effect of sea-floor oxygenation impairs depth reconstructions, we studied the percentage of

planktonic foraminifera (%P) in five well-dated sedimentary successions from the Lower Pliocene of Crete, Corfu and

Milos in Greece. Additionally, we assessed whether different foraminiferal size fractions and counting methods affect the

determination of the percentage of planktonic foraminifera. The palaeobathymetric evolution calculated for each basin was

confirmed for all successions by an independent check on depth-related occurrences of benthic foraminifera. After

correction for bathymetry changes of the basin due to sedimentation, compaction and eustatic sea level variations, the

0031-0182/$ - s

doi:10.1016/j.pa

T Correspondi

116 2523629; fa

E-mail addr

laeoecology 221 (2005) 245–265

ee front matter D 2005 Elsevier B.V. All rights reserved.

laeo.2005.02.013

ng author. Present address: Geology Department, University of Leicester, University Road, Leicester LE1 7RH, UK. Tel.: +44

x: +44 116 2523918.

ess: [email protected] (D.J.J. van Hinsbergen).

Page 2: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265246

vertical movement history of the basin floor was inferred. We propose a standard methodology for reconstructions of

palaeobathymetry of marine sedimentary successions from foraminiferal associations.

D 2005 Elsevier B.V. All rights reserved.

Keywords: Milankovitch cyclicity; Eccentricity; (Palaeo-)bathymetry; P/B ratio; Vertical movements; Greece

1. Introduction

One way to determine the timing and rates of

vertical movements of the basin floor during sed-

imentation is to reconstruct variations in palaeodepth

from the basin’s sediments, together with the sediment

accumulation history (geohistory analysis: Van Hinte,

1978). Palaeobathymetry estimates can be further

derived from faunal distribution patterns. To this

end, foraminifera are useful tools. Bandy (1953) and

Bandy and Arnal (1960) already outlined the extra

information that can be derived from depth distribu-

tions of benthic foraminifera when reconstructions of

basin configurations and vertical movements are

attempted, although no detailed quantitative estimates

were made about subsidence and uplift. However, as

Van Hinte (1978) stated referring to microfossils in

general: bPalaeo-water-depth determination is a

highly complex artQ. Information on depth distribu-

tions of fossil benthic foraminifera is usually obtained

from indirect evidence, or by comparison with living

representatives of the taxa present, or by comparing

functional morphology of living and fossil taxa. A

complicating factor is that depth distributions of

Recent benthic foraminifera may differ between

basins, as was demonstrated by Parker (1958) and

Bandy and Chierici (1966), although the latter authors

found a number of species to be isobathyal. Moreover,

some taxa appear to be able shift to different depth

zones in different environmental conditions (e.g.

Pflum and Frerichs, 1976; Speijer et al., 1997; De

Rijk et al., 2000). Although no consensus exists as yet

about depth distributions of fossil benthic foramin-

ifera, certain taxa are often used to constrain palae-

odepths within limits of several hundreds of meters,

and by using combinations of taxa occurring in a

sample the palaeodepth can be more precisely con-

strained, provided that resedimentation and reworking

can be excluded. However, this method asks for more

than superficial knowledge of benthic foraminifera.

Another option mentioned by Van Hinte (1978)

was to use the ratio between planktonic and benthic

foraminifera (P/B ratio) as a tool to reconstruct

palaeodepths. The abundance of benthic foraminifera

generally reaches a maximum on the outer shelf and

the upper slope. Planktonic foraminifera are virtually

absent in neritic environments and their abundance

generally increases with increasing water depth. The

ratio, expressed as the percentage of planktonic

foraminifera (%P), increases with water depth and

distance to shore to reach nearly 100% in lower

bathyal and abyssal environments (Douglas and

Woodruff, 1981; Berger and Diester-Haas, 1988).

Theoretically, the %P would thus seem directly

related to water depth, but this is not always the case.

Studies on Recent material and in laboratory experi-

ments have shown that a strong relationship exists

between the oxygen level of bottom waters and the

abundance and diversity of benthic foraminiferal

populations (e.g. Harman, 1964; Sen-Gupta and

Machain-Castillo, 1993; Kaiho, 1994, 1999; Loubere,

1994, 1996, 1997; Alve and Bernhard, 1995; Jorissen

et al., 1995; McCorkle et al., 1997; Moodley et al.,

1998; Jorissen and Wittling, 1999). These findings,

when applied to the fossil record, are confirmed by

sedimentological and geochemical evidence (e.g.

Nolet and Corliss, 1990; Rohling et al., 1993;

Nijenhuis et al., 1996; Den Dulk et al., 1998; Jorissen,

1999a,b; Seidenkrantz et al., 2000). As the oxygen

level of bottom waters decreases–for instance through

increased primary productivity in surface waters and

the consequent increased input of organic matter–the

abundance of benthic foraminifera will initially

increase and then rapidly decrease until benthic life

is no longer possible (e.g. Verhallen, 1991; Jorissen,

1999b). Hence, the %P is not only determined by

depth, but also by changes in oxygenation state of the

bottom waters. The concentration of oxygen in bottom

waters in turn may fluctuate under the influence of

astronomically induced climate variations, influencing

Page 3: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 247

bottom water ventilation (e.g. De Visser et al., 1989;

Hilgen et al., 1995; Lourens et al., 1996; Kouwen-

hoven et al., 2003). To us, longer-term variations

(eccentricity: 100 and 400 ky) are of special interest,

since on this time scale tectonically induced vertical

movements of the basin floor can also play a

significant role.

Van der Zwaan et al. (1990) determined a

regression for the relationship between bathymetry

and the percentage of planktonic foraminifera with

respect to the total fossil foraminiferal population

(%P), based on present-day bathymetric transects:

Depth mð Þ ¼ e3:58718þ 0:035344%Pð Þ ð1Þ

where %P=percentage planktonics in the total fora-

miniferal association, calculated as 100*P/(P +B),

P=number of planktonic specimens and B =number

of benthic specimens.

In their calculations, Van der Zwaan et al. (1990)

discarded a number of species from the benthic

population, because these were considered to be deep

Corfu

20 22 2434

36

38

40

42

Milos

Crete

GREECE

ALBANIA

F.Y.R.O.M.BU

*

Akrotira

N

AEGE

Corfu Coast

Tsouvala

0 50 100

km

Fig. 1. Map of the Aegean are

infaunal (i.e. dwelling well below the sediment–water

interface) and therefore not directly dependent on the

flux of organic matter to the sea floor, which forms the

basis for the depth relation of regression (1) (Suess,

1980; Berger and Diester-Haas, 1988; Van der Zwaan

et al., 1990). These infaunal species discarded from

the regression calculations were the benthic genera

Bulimina, Bolivina, Globobulimina, Uvigerina and

Fursenkoina. Several of these taxa are seen to

dominate benthic assemblages under unfavourable

circumstances and are then commonly indicated as

stress-markers. These should be omitted from the

determination of %P. The plankton fraction %P is

then calculated as

%P ¼ 1004 P= P þ B� Sð Þð

where S=number of stress markers (deep infauna).

Regression (1) was constructed from modern

transects in the Gulf of Mexico, the Gulf of California,

the west coast of the USA and the Adriatic Sea,

yielding near-identical results. Values of %P of 0 and

26 28

LGARIA

TURKEY

***

**

ki

AN

SEA

Kalithea

Aghios Vlassios

a with sample locations.

Page 4: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265248

100 lead to calculated depths of 36 and 1238 m,

respectively. The standard error increases with

increasing %P, and in the original regression of Van

der Zwaan et al. (1990), the 90% confidence limit at a

single %P value of 50% (430 m) was approximately

100–150 m, and at a %P value of 99% (1200 m) 400

m. The first and most important way to improve the

resolution of the depth estimate is to use large

numbers of samples covering a small time interval.

Additionally, Van der Zwaan et al. (1990) noted that

thorough screening of the samples for resedimentation

also significantly adds to a higher resolution of the

bathymetry estimate.

To determine the influence of the amount of

oxygen in bottom waters on %P (and to distinguish

between this effect–if present–and bathymetry varia-

tion), we studied the foraminiferal content of five

well-dated sedimentary successions in the Lower

Pliocene of Crete, Milos and Corfu (Greece) (Fig.

1). Various authors have used regression (1) of Van

der Zwaan et al. (1990) to reconstruct tectonic vertical

movements in sedimentary basins (e.g. Barbieri,

1992; Meulenkamp et al., 1994; Grafe, 1999; Van

der Meulen et al., 1999, 2000; Ten Veen and

Kleinspehn, 2000; Baldi et al., 2001; Van Hinsbergen

et al., 2004) and every paper makes different choices

regarding omission of samples and species and the

assumptions on the resolution of the method. There-

fore, we propose a standard methodology for the

reconstruction of palaeodepths based on foraminiferal

associations and apply this to reconstruction of

vertical movements of the basin floor.

2. Geologic setting and successions

The selected sedimentary successions on Corfu,

Crete and Milos each occupy a distinct structural

position within the Hellenides (Fig. 1). During the

Early Pliocene, Corfu was situated in a foreland basin,

and characterised by a high sedimentation rate and

deposition of clays and sandy turbidites (Linssen,

1991; Weltje and De Boer, 1993). The succession at

Corfu Coast is located along the northwestern coast of

Corfu, where approximately 500 m of clays and sandy

turbidites were sampled at 100 levels at 5 m intervals.

Linssen (1991) magneto- and biostratigraphically

dated the succession (Fig. 2). Milos was situated in a

restricted back-arc basin that developed in the central

part of the Aegean (Fig. 1). The successions near

Tsouvala and Akrotiraki were sampled in the south of

Milos (for exact locations, see Van Hinsbergen et al.,

2004) and contain 20–30 m of marl-sapropel alter-

nations. The Akrotiraki succession was sampled at 99

levels and the Tsouvala succession at 95 levels. On

central Crete, Early Pliocene sedimentation occurred

in a narrow graben cross-cutting the Aegean arc

(Meulenkamp, 1985; Meulenkamp et al., 1994). The

successions near Aghios Vlassios and Kalithea were

sampled in the central northern part of Crete (Figs. 1

and 2; for detailed locations and descriptions see

Spaak, 1983; Jonkers, 1984; Driever, 1988). These

successions consist of alternating marls and sapropels

and were sampled at 141 and 162 levels, respectively.

Previously, micropalaeontological analyses were car-

ried out by Spaak (1983), Jonkers (1984) and Driever

(1988), and Bianchi et al. (1985) dated some volcanic

ash layers. This age information was used to correlate

the cyclic alternations of sapropels and marls to the

astronomical polarity time scale (APTS: Hilgen, 1991;

Fig. 2). The Cretan successions were used for a vertical

movement study by Meulenkamp et al. (1994).

3. Screening of the successions and samples

3.1. Age control

In order to assess the effect of orbitally induced

trends, the successions should preferably represent at

least 400 ky–a full eccentricity cycle–and should be

well dated, preferentially by tuning to the astronomical

polarity time scale (e.g. Hilgen, 1991; Krijgsman et al.,

1995; Sierro et al., 2001; Abdul Aziz et al., 2003).

Moreover, to increase the accuracy of palaeobathyme-

try and to reconstruct long-term eccentricity induced

trends, one sample per 10–20 ky is preferentially

analysed. The successions on Crete are well dated by

biostratigraphy, can be astronomically tuned, and have

a sample spacing of 1 sample per 10–15 ky (Spaak,

1983; Jonkers, 1984; Driever, 1988). The Corfu Coast

succession is accurately dated by magnetostratigraphy

(Linssen, 1991), with a sample spacing of 1 per 5 ky.

The successions on Milos have a sample spacing of

10–15 ky and are dated by bio-, cyclo- and magneto-

stratigraphy (Van Hinsbergen et al., 2004).

Page 5: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

marl breccia

- H -

Tsouvala

20

25

30

Akrotiraki

0 m

5

10

15

20

25

30 . . . . . . .. . . . . . .

. . . . . . .. . . . . . .

. . . . . . . .. . . . . . .

(Mi los) (Mi los)

- H -

insolation

Aghios Vlassios (Crete)

400 500 600 W/m

Corfu Coast (Corfu)

0 m

100

200

300

400

500

Kalithea (Crete)

0

AB

C

E

F

H

F

C

G

E

KEY. . . .. . .

alternating marlsand sapropels

marl

volcanic ash

diatomaceous marl

pyroclastic deposits

sand

alternating marlsand turbiditic sandstones

5.5 Ma

5.0 Ma

4.5 Ma

4.0 Ma

3.5 Ma

3.0 Ma

50

100

50

15

10

5

0 m

5.5 Ma

5.0 Ma

4.5 Ma

4.0 Ma

3.5 Ma

3.0 Ma

2

. . . .

- H - hiatus

unconformity

Magnetic polaritytimescale

normal magnetic polarity

reversed magnetic polarity

J

D

C

Fig. 2. Bio-, magneto-, and cyclostratigraphic correlations between the successions of Milos, Crete and Corfu. The cyclic successions are tuned

using the target curve of Laskar et al. (1993). For biostratigraphy of the Cretan successions, see Spaak (1983), Jonkers (1984) and Driever

(1988), cyclostratigraphic correlation of these successions is further explained in Van Hinsbergen and Meulenkamp (submitted for publication).

Bio- and cyclostratigraphy of the Milos successions is carried out by and explained in Van Hinsbergen et al. (2004). Magnetostratigraphic dating

of succession Corfu Coast was carried out by Linssen (1991). Ages of the bioevents are taken from Lourens et al. (2004): A=First Occurrence

(FO) Globorotalia margaritae; B=Last Occurrence (LO) Reticulophenestra antarctica (4.91 Ma); C=FO G. puncticulata (4.52 Ma); D=First

Common Occurrence (FCO) Gephyrocapsa spp. (4.33 Ma); E=FCO Discoaster asymmetricus (4.12 Ma); F=Last Common Occurrence (LCO)

G. margaritae (3.98 Ma); G=LCO Sphenolithus spp. (3.70 Ma); H=LO G. puncticulata (3.57 Ma); J=FO G. bononiensis (3.31 Ma).

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 249

Page 6: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

Plate I. Depth markers. (see page 251)

1. Ammonia beccarii;

2. Discorbis sp.;

3. Elphidium macellum;

4. Elphidium gerthi;

5. Cibicides lobatulus;

6. Cibicides ungerianus;

7. Cibicides pseudoungerianus;

8. Gyroidina soldanii;

8. Cibicides pachydermus;

9. Uvigerina peregrina;

10 and 11. Uvigerina peregrina;

12. Uvigerina proboscidea;

13. Uvigerina hispida;

14. Uvigerina semiornata rutila;

15. Planulina ariminensis;

16. Siphonina reticulata. SEM pictures from: 1: Jorissen (1988); 2, 5, 6, 7, 8, 10, 11, 12, 13: Den Dulk (2000); 3: http://

www.ucl.ac.uk/GeolSci/micropal; 4: http://palaeo.electronica.org/2002_2/guide/rota.htm; 9, 14, 15, 16: Kouwenhoven (2000);

scale bar 1 Am.

Plate II. Depth markers. (see page 252)

1. Oridorsalis stellatus;

2. Cibicides kullenbergi;

3. Cibicides bradyi;

4. Cibicides robertsonianus;

5. Karreriella bradyi;

6. Eggerella bradyi;

7. Cibicides wuellerstorfi;

8. Cibicides italicus. SEM pictures from: 1, 3, 4, 8: Kouwenhoven (2000); 2, 5, 6, 7: Den Dulk (2000); Scale bar 1 Am.

Plate III. Stress markers. (see page 253)

1. Cancris auricula;

2. Valvulineria bradyana;

3. Bolivina striatula;

4. Bolivina spathulata;

5. Bolivina alata;

6. Bolivina tortuosa;

7. Bolivina dilatata;

8. Chilostomella oolina;

9. Bulimina marginata;

10. Bulimina aculeata, marginata type;

11. Bulimina aculeate;

12. Bulimina aculeate;

13. Bulimina exilis;

14. Fursenkoina pauciloculata;

15. Stainforthia fusiformis;

16. Globobulimina spp.;

17. Hopkinsina (Uvigerina) pacifica;

18. Uvigerina (Rectuvigerina) cylindrica cylindrica;

19. Uvigerina (Rectuvigerina) cylindrica gaudryinoides;

20. Bolivina plicatella;

21. Bulimina alazanensis. SEM pictures from 1, 3, 7, 9, 14, 15, 17: Barmawidjaja (1991); 2, 10, 12: Jorissen (1988); 4, 5, 8, 11, 13,

16, 21: Den Dulk (2000); 6, 18, 19, 20: Kouwenhoven (2000); Scale bar 1 Am.

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265250

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Plate I (caption on page 250).

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 251

Page 8: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

Plate II (caption on page 250).

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265252

Page 9: Paleobathymetry in the backstripping procedure: …forth/publications/Hinsbergen_2005c.pdfderived from faunal distribution patterns. To this end, foraminifera are useful tools. Bandy

Plate III (caption on page 250).

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 253

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1

2

3

4

5

6

7

8

9

10

11

12

13

Fig. 3. Schematic drawing of a picking tray. In grey the fields of the

counting scheme we applied are indicated, with numbers indicating

the counting order.

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265254

3.2. Suitability of samples

Most suitable for palaeobathymetry reconstruc-

tions are fine-grained pelagic marine sediments that

were deposited in low-energy environments. These

sediments generally contain faunal populations that

are least affected by downslope transport. Bandy

(1953), referring to Natland and Kuenen (1951)

already stated: bone might expect coarser laminae

representing turbidity flows, etc., to carry the

B

0

20

40

60

80

100

%P

(sp

read

2)

0

20

40

60

80

100

%P

(sp

read

3)

%P (sp0 20 40

%P (spread 1)0 20 40 60 80 100

0

20

40

60

80

100

%P

(63

-125

)

A

0

20

40

60

80

100

%P

(20

0-59

5)

%P (120 20 40

%P (125-595)0 20 40 60 80 100

Fig. 4. A) Cross-plots showing reproducibility of %P counts. Difference

counts used to construct this figure are given in online Appendix I. B) Cros

than the 125–595 Am fraction used by Van der Zwaan et al. (1990) gives s

are given in online Appendix II.

shallow-water species; whereas the thin shales,

representing gradual deposition, would carry the

deep-water populationsQ.Individual samples are suitable for analysis of the

%P if:

– No dissolution of carbonate has occurred. Dis-

solution of carbonate will preferentially affect

planktonic, rather than benthic foraminifera (Van

der Zwaan et al., 1990; Boltovskoy and Totah,

1992), thus influencing the %P. Samples with

evidence for carbonate dissolution, also evidenced

by abundant fragmented shells, should be dis-

carded. Our criterion was, that preservation should

allow determination of benthic foraminifera at

species level.

– The sample is not contaminated by down-slope

transport. To determine whether the sample was

likely to contain transported foraminifera, we

checked for presence of quartz grains or rock

fragments in the washed and sieved fraction of the

sample, and for benthic taxa expected to live at

very different water depths (Bandy, 1953). In

%P (spread 1)0 20 40 60 80 100

0

20

40

60

80

100

%P

(sp

lit)

read 1)60 80 100

5-595)60 80 100

s between the preading and splitting procedures are minimal. The

s-plots, showing that a smaller or larger size fraction of foraminifera

ignificantly different results. The counts used to construct this figure

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5.5 5.0 4.5 4.0 3.5 3.0Age (Ma)

0

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

100

%P

%P

%P

%P

%P

Fig. 5. Plots of the percentage of planktonic foraminifera (%P)

versus age of the successions. The counts used to construct this

figure are given in online Appendix III.

0

20

40

60

80

100

3.54.04.55.05.5

~110 kyr (9-point) moving average

~37 kyr (3-point) moving average

Age (Ma)

%P

Fig. 6. Graph of the results of the Upper Pliocene Kalithea

succession on Crete (Greece), illustrating the visual effect of a

moving average filter.

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 255

combination with broken specimens and/or a size

sorting of planktonic and benthic foraminifera, this

might indicate transportation and/or winnowing.

– Finally, we omitted all samples from sapropels,

since these were deposited during more or less

severe anoxic conditions, which significantly

modifies %P values (see above).

Samples that passed these criteria were considered

suitable for analysis of %P. All samples of the

Aghios Vlassios and Kalithea successions, together

with 53 samples of the Corfu Coast succession, 57

samples of the Tsouvala succession and 77 samples

of the Akrotiraki succession were used for further

analysis.

4. Analysis of the percentage of planktonic

foraminifera (%P) and the percentage of stress

markers (%S)

4.1. Basic taxonomic concept

Some basic taxonomic knowledge about foramin-

ifera is needed for the analysis of the %P. For images

of planktonic and benthic foraminifera, and in order to

discriminate between taxonomic groups at a generic

level, the reader is referred to e.g. Loeblich and

Tappan (1964, 1988). Accessible taxonomy on the

species level can be found in, for instance, AGIP

(1982) and Jones (1994). For convenience, some

common Neogene taxa are shown in Plates I–III.

As above mentioned, %P depends not only on

depth, but also on the oxygen level of bottom waters.

When oxygen levels are low, benthic associations will

change and genera and species with high tolerance to

stressed conditions tend to flourish. Under normal,

oxic conditions, these genera will generally have an

infaunal habitat. In many cases the infaunal genera

that were discarded from the %P determination by

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D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265256

Van der Zwaan et al. (1990) can be considered as

stress markers, especially when they occur in rela-

tively high numbers. We propose to calculate the

percentage of stress markers with respect to the total

benthic foraminiferal population (%S) separately. We

have included recent progress on knowledge about

benthic foraminifera in stressed environments in our

list of stress markers, modifying the original list of

Van der Zwaan et al. (1990) to include: Bolivina spp.,

except for B. plicatella and B. pseudoplicata, non-

costate Bulimina, Uvigerina spp., except for U.

5.5 5.0 4.5 4.0 3.5 5.5Age (Ma)

Corfu Coast

Milos-Tsouvala

Milos-Akrotiraki

Crete-Aghios Vlassios

0

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

100

%P

%P

%P

%P

%P

Crete-Kalithea

3.0

5.5 5.0 4.5 4.0 3.5 5.5Age (Ma)

3.00.000.020.04

Ecc

entr

icity

Var

iatio

n

Fig. 7. Approximately 100 ky moving average curves of the percentage o

among the total benthic population (%S) versus time. Grey interval around

averaged interval. The curves are correlated to the Eccentricity La93 curve

4.8 and 4.4 Ma are easily recognised in both the %S and %P diagrams. The

on the curves.

semiornata, Rectuvigerina spp., Valvulineria spp.,

Cancris spp., Fursenkoina spp., Stainforthia spp.,

Globobulimina spp. and Chilostomella spp. (Seiglie,

1968; Van der Zwaan, 1982; Jonkers, 1984; Van der

Zwaan et al., 1985, 1999; Van der Zwaan and

Jorissen, 1991; Verhallen, 1991; Rathburn and Cor-

liss, 1994; Fariduddin and Loubere, 1997; Jorissen,

1999b; Kouwenhoven, 2000) (Plate I).

In summary, we counted planktonic and benthic

foraminifera, the latter subdivided in dnormalT open

marine taxa and stress markers.

Crete-Kalithea

5.0 4.5 4.0 3.5 3.0Age (Ma)

Corfu Coast

Milos-Tsouvala

Milos-Akrotiraki

Crete-Aghios Vlassios

0

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

0/100

20

40

60

80

100

%S

%S

%S

%S

%S

5.0 4.5 4.0 3.5 3.0Age (Ma)

0.000.02

0.04

Eccentricity

Variation

f planktonic foraminifera (%P) and the percentage of stress markers

the %P curve represents the standard deviation on the depth of the

(Laskar et al., 1993). The well-defined eccentricity minima around

less pronounced minima around 4.0 and 3.6 Ma have less influence

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0

20

40

60

80

100

%P

and

% S

3.54.04.55.05.5Age (Ma)

%S%P (not corrected for %S)%P (corrected for %S)

Fig. 8. Graph showing the effect of the correction for stress markers

on %P. Correction for stress markers will diminish the amplitude of

%P oscillations resulting from oxygenation variations, but %P

variation still remains. The intervals of least oxygen induced stress–

corresponding to the intervals of minimal %S–are the most reliable

to estimate the true palaeobathymetry form. For the astronomically

tuned eustatic sea level curve since 15 Ma of Lourens and Hilgen

(1997), see online Appendix IV.

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 257

4.2. Counting procedures

4.2.1. Split versus spread

To obtain a representative collection of foramin-

ifera, samples can repeatedly be split in half, until the

desired number of specimens is reached. This method

was used by e.g. Jonkers (1984), who counted splits

of samples of the Kalithea and Aghios Vlassios

successions until 200 benthic specimens were

obtained. This method, however, is rather time-

consuming.

We applied a second, much quicker method,

which starts with putting approximately half of the

sample in one corner of the picking tray and then

randomly spreading the required amount on the rest

of the tray. This method corrects for size fractionation

when the sample is strewed. Our counting was

carried out following the predetermined scheme

shown in Fig. 3. As a rule, a minimum of 100

specimens of the larger group, benthic or planktonic

foraminifera, was counted.

To evaluate whether the results obtained by both

methods are statistically different, 30 samples of the

Kalithea succession, splits of which were previously

counted by Jonkers (1984), were spread and counted

three times by two different people using the spread-

ing procedure. The cross-plots of Fig. 4A show that

the results are reproducible, using either way of

counting.

4.2.2. Influence of the size of the sieved fraction on

%P

Jonkers (1984) provided counts of three different

size fractions, based on 54 samples from the Kalithea

succession. A clear difference in %P exists between

the 125–595 Am fraction and the smaller (63–125 Am)

and larger (200–595 Am) fraction (Fig. 4B). For the

construction of the regression (1), the fraction of 125–

595 Am was used (Van der Zwaan et al., 1990), which

for reasons of comparison should thus be considered

as the standard for our palaeobathymetric analysis.

4.3. Results

We determined the %P and %S of the successions

at Crete, Milos and Corfu following the procedures

described in the previous sections. The counts were

obtained by spreading the 125–595 Am fraction on a

picking tray and counting according to the scheme of

(Fig. 3), with a minimum of 100 specimens of either

planktonic or benthic foraminifera. The results are

shown in Fig. 5. The age is constrained by bio-,

magneto- and cyclostratigraphy. Fig. 5 will be used as

the basis for further interpretation and discussion.

5. Interpretation of the results and discussion

The diagrams of Fig. 5 reflect the interplay

between the effect on %P of oxygen level of the

bottom waters and depth variation. We aim to

discern between palaeobathymetry-induced and oxy-

genation-induced variations of %P. High-amplitude

short-term oscillations of the %P (b100 ky, e.g.

across sapropels: Jorissen, 1999b; Kouwenhoven,

2000) are unlikely to result from true palaeobathy-

metry variations. To filter the long-term %P trend,

which is more likely to result from both palae-

obathymetry and oxygenation variations, one can use

a moving average, which for the Kalithea succession

is illustrated in Fig. 6. In Fig. 7 the %P curves of the

five successions are plotted, applying a moving

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5.5 5.0 4.5 4.0 3.5Age (Ma)

+200

0

-200

-400

-600

-800

-1000

-1200/+200

0

-200

-400

-600

-800

-1000

-1200/+200

0

-200

-400

-600

-800

-1000

-1200/+200

0

-200

-400

-600

-800

-1000

-1200/+2000

-200

-400

-600

-800

-1000

-1200

sea level

sedimentaccumulation

paleobathymetry

5.5 5.0 4.5 4.0 3.5

Age (Ma)3.0

-200

-400

-600

-800

-1000

Sub

side

nce

0

0

-200

-400

-600

-800

-1000

-200

5.5 5.0 4.5 4.0 3.5Age (Ma)

3.0

5.5 5.0 4.5 4.0 3.5

Age (Ma)3.0

0

200

400

600

800

=vertical motion =oxygen effect

A. Geohistory B. Vertical motionU

plift

Sub

side

nce

Uplift

Sub

side

nce

Uplift

dept

h / t

hick

ness

dept

h / t

hick

ness

dept

h / t

hick

ness

dept

h / t

hick

ness

dept

h / t

hick

ness

}}

Fig. 9. Construction of vertical movement curves obtained by correction of the palaeobathymetry for sedimentary infill, compaction and eustatic

sea level fluctuations. The sea level curve is taken from Lourens and Hilgen (1997) (see Appendix II). The base of the oldest succession is taken

as reference level and thus set at 0 m. As a result, Corfu and Milos reveal subsidence, and a negative vertical axis, whereas Crete shows uplift

and a positive vertical axis.

Fig. 10. A. Depth distribution of a selection of benthic marker species, based on Parker (1958), Blanc-Vernet (1969), Wright (1978), Parisi

(1981), Jorissen (1987), Sprovieri and Hasegawa (1990), Sgarella and Moncharmont Zei (1993), De Stigter et al. (1998), De Rijk et al. (2000),

Seidenkrantz et al. (2000), Jannink (2001), Kouwenhoven et al. (2003). Note that these studies are all based on the Mediterranean. See Plates II

and III for images of these species. B. Results of the taxonomic check for a selection of samples of the various successions. The depth values

calculated from the %P are shown in the left-hand column and the depth range estimated from the depth marker species of (A) are represented as

horizontal bars. The results confirm the observed depth trends, although the depth estimated for the Kalithea succession based on the taxa is

somewhat shallower than the calculated depth, whereas the calculated depth of the Aghios Vlassios succession is shallower than estimated by

benthic taxa.

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265258

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100 200 500 750 1000 m0

sam

ple

calc

ula

ted

d

epth

GR 3260 - 486 m

GR 3300 - 321 m

GR 3316 - 229 m

Ag

hio

s V

lass

ios

(Cre

te)

Akr

oth

irak

i (M

ilos)

GR 10504 - 579 m

GR 10494 - 860 m

GR 10463 - 402 m

GR 10454 - 619 m

Co

rfu

Co

ast

GR 11382 - 1134 m

GR 11415 - 981 m

GR 11436 - 853 m

GR 11442 - 939 m

GR 11455 - 893 m

GR 11477 - 875 m

Kal

ith

ea (

Cre

te)

GR 2665 - 864 m

GR 2737 - 570 m

GR 2209 - 418 m

GR 2225 - 581 m

GR 2243 - 267 m

GR 2700 - 693 m

B

diversity highhighintermediate low very lowintermediate

Ammonia spp.

epiphytes

Elphidium spp.

Cibicides pachyderma

Uvigerina spp.

C. (pseudo)ungerianus

Planulina ariminensis

C. bradyi/robertsonianus

Cibicidoides kullenbergi

Oridorsalis spp.

Eggerella/Karreriella spp.

C. wuellerstorfi / C. italicus

Gyroidina spp.

Siphonina reticulata

Cibicides lobatulus

depth 100 200 500 750 1000 m0Aag

e (M

a)

4.601

4.720

4.782

4.835

4.969

5.083

4.216

4.374

4.678

4.835

3.704

3.919

4.080

4.353

4.860

5.244

3.215

3.300

3.570

CP 2364 - 830 m4.388

CP 2264 - 427 m3.745

CP 2229 - 713 m4.056

palaeodepth estimate based on marker species

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 259

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D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265260

average that approximates 100 ky (50 ky before and

after each data point).

The %P curves, constructed from the successions

from Milos and Crete reveal a cyclic fluctuation of

%P, superimposed on a decreasing and increasing

trend on Crete and Milos, respectively. This cyclicity

corresponds to the 400 ky eccentricity-driven inso-

lation fluctuation, shown in the curves of Laskar

(1990) and Laskar et al. (1993) (Fig. 7). Fig. 7 also

includes the curves of the percentage stress markers

among the total benthic population (%S) in these

successions, showing, that the cyclic decrease and

increase in %P corresponds to an increase and

decrease in %S, respectively. This suggests that the

400 ky fluctuations are caused by changes in

bottom-water oxygenation. If one aims to reconstruct

the bathymetry variations, data points representing

comparable oxygen levels should be connected,

preferably the best-ventilated intervals, as regression

(1) was constructed on the basis of samples taken

from well-ventilated intervals and disregarding

infauna, largely corresponding to our stress markers

(Van der Zwaan et al., 1990). These intervals can be

recognised, because they coincide with the intervals of

lowest %S (Figs. 7 and 8).

It should be noted that the oxygenation variations

not only influence the %S, but also the %P: abundance

of the total benthic population appears to fluctuate with

oxygenation variations, assuming that primary pro-

duction of planktonic foraminifera is constant (Fig. 8).

Correction for stress markers will therefore diminish

the amplitude of the oxygenation-induced %P varia-

tion, but still oxygenation-induced variations remain in

the %P-curves. This explains the strong 400 ky %P

variations in the curves of Figs. 7–9, despite the fact

that stress markers are corrected for. The omission of

stress markers from the benthic population therefore

will not fully correct for oxygenation effects on

benthic populations, but measuring %S provides a

strong tool to distinguish between oxygenation and

depth-effects on the %P, as the %S is not directly

related to water depth. Therefore, %P variations

resulting from oxygenation variation will be accom-

panied by %S variations, whereas %P variations as a

response to true depth changes will not be.

Cyclic fluctuations on Milankovitch time-scales

are absent in both the %P and %S curves constructed

from the Corfu Coast succession, which is in line with

its position in a wide and deep, and therefore probably

well-ventilated foreland basin. Restricted basins, for

instance silled basins are generally less well venti-

lated, as was already noted by Bandy (1953) (See also

Krijgsman, 2002). The high-resolution %P and %S

curves obtained from the successions of Milos and

Crete allow us to discern unambiguously between

oxygen-induced and bathymetry-induced effects on

the %P curve. If palaeobathymetry-analysis is attemp-

ted on successions with a poorer time resolution, this

distinction can be more difficult or even impossible to

make. In such cases, samples with very low %S levels

probably best represent the true bathymetry, whereas

samples with increasing %S give a decreasing

confidence in the depth estimate. A case study on

the Kalithea succession shows, that if all samples with

%S N60 are discarded the closest match is made

between the constructed and the best-ventilated depth

trend. Discarding more samples (e.g. %S N40 or 50)

will not yield a better match, but it will decrease the

amount of data drastically. It should be noted that in

cases of lower-resolution successions the bathymetry

estimation has a much larger uncertainty and should

be interpreted accordingly.

In summary, the %P trend is strongly influenced

by oxygen depletion in poorly ventilated basins and

the most reliable bathymetry trend can be constructed

from the dbest-ventilatedT intervals. It is now possible

to calculate the bathymetry from the %P curves by

using regression (1).

The resulting moving average curves reveal a

shallowing trend on Corfu and Crete, and a deepening

trend on Milos (Fig. 9).

6. Taxonomic check

Any significant trends in palaeobathymetry can be

independently checked on selected samples along the

succession, by identifying marker species for

selected depth intervals. Fig. 10A and Plates II and

III show a selection of benthic species from Middle

Miocene to Recent that can well be used as depth

markers. One should keep in mind, however, that

many of these taxa are in themselves sensitive to

low-oxygen conditions on the sea floor. Unexpected

absence of these depth markers may be related to

environmental conditions as well as to depositional

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Sample

D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265 261

depth. The depth markers confirm the depth trends

calculated from the %P (Fig. 10B).

Can species be recognised?

yes

noDISCARD SAMPLE

Evidence fordownslope transport

no

yesDISCARD SAMPLE

Evidence forreworking?

no

yesDISCARD SAMPLE

Determine %P and %S

Construct curves;choose moving average

Cyclic fluctuationin %P?

yes

no

Matching cyclic fluctuation in %S?

no

Select points withlowest %S

Calculate depth (D= e )Construct paleobathymetry curve

Confirmation from depth markers

-add thickness (m) of accumulated sediment-add amount of compaction (m)-correct for eustatic sea level changes-correct for flexural effects

Construct vertical movement curve

Interpret trends in vertical movement

yes

%P=P/(P+B-S)*100%S=(S/B)*100

3.58718+(0.03534*%P)

BACKSTRIPPING:

Fig. 11. Flow chart, showing the subsequent steps from sampling o

the succession toward an interpretable vertical movement curve

P=number of specimens of planktonic foraminifera, B =number o

specimens of benthic foraminiferal, S =number of specimens o

stress markers (see text for further explanation).

7. From palaeobathymetry to vertical movement

To reconstruct vertical movements from these

palaeobathymetry trends, corrections should be car-

ried out, which have been described in backstripping

procedures of e.g. Steckler and Watts (1978), Watts et

al. (1982) and Steckler et al. (1999). Sediment

deposition in a basin will have a shallowing effect.

To construct the true movement of a chosen reference

level (normally, the first sample level of the succes-

sion), the thickness of the accumulated sediment

should be added to the estimated palaeobathymetry

at each datum. Additional corrections can be carried

out for compaction of the sediment. Van Hinte (1978)

and Van der Meulen et al. (1999) used the present

thickness–initial thickness relationships for sedimen-

tary rocks of Perrier and Quiblier (1974) to correct for

compaction. This relationship shows a general reduc-

tion of the sedimentary column of 15–25%, but this is

strongly determined by lithology and amount of

overburden. Not correcting will therefore yield an

overestimation of subsidence or an underestimation of

uplift. The present thickness of succession Corfu

Coast is 500 m, which, following Perrier and Quiblier

(1974) must originally have been approximately 600

m assuming that no large overburden was once

present on top of Corfu Coast. On Milos and Crete,

the amount of compaction must have been of the order

of 10–20 m and hence is neglected.

Finally, to determine the true movement of the

reference level, bathymetry variation due to eustatic

sea level changes should be corrected for. An

astronomically tuned eustatic sea level curve since

the Middle Miocene with respect to the present-day

sea level was constructed by Lourens and Hilgen

(1997). Fig. 9b shows the resulting vertical movement

curves for southern Milos, the north of Central Crete

and northeastern Corfu. In case of the Milos and

Cretan successions, the last corrections have only

minor influence and the deepening and shallowing

trends are caused by subsidence and uplift, respec-

tively (Fig. 9). The curve obtained from the Corfu

Coast succession indicates approximately 200 m of

shallowing, but reveals several hundreds of meters of

f

.

f

f

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D.J.J. van Hinsbergen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 245–265262

subsidence after correction for sedimentation and

eustatic sea level changes (Fig. 9). The older

succession Akrotiraki shows evidence for subsidence

between 4.8 and 4.4 Ma, whereas the younger

succession Tsouvala reveals slight uplift between 4.4

and 4.0 Ma (Figs. 9 and 10). The resulting movement

curves are the combined effect of tectonic motions

and isostasy (e.g. due to sediment loading or crustal

thickening/thinning).

8. From the percentage of plankton to vertical

movements: a flow chart

The procedures followed in the determination of

the vertical movements in the five successions

discussed in this paper are summarised in a flow

chart (Fig. 11). In general, the steps in the flowchart

will lead to an accurate reconstruction of vertical

movements. In practice, however, successions might

not contain only fine clay, may not be accurately dated

or may comprise only little time spans. Generally, the

accuracy of the palaeobathymetry reconstruction

should be discussed carefully.

Acknowledgements

We acknowledge the valuable comments of Finn

Surlyk and incisive reviews by Hans Thierstein and an

anonymous reviewer. Additionally Francisco Sierro is

thanked for comments on an earlier version of the

manuscript. Counting of the Kalithea and Aghios

Vlassios samples was carried out by Rudi de Koning.

Gerrit van ’t Veld and Geert Ittman are thanked for the

preparation of the samples. Samples of the Corfu

Coast succession were kindly provided by Cor

Langereis of the Palaeomagnetic Laboratory dFortHoofddijkT. Rinus Wortel, Johan Meulenkamp, Rein-

oud Vissers and Bart Meijninger are thanked for

critical reading of the manuscript.

Appendix A. Supplementary data

Supplementary data associated with this article can

be found, in the online version, at doi:10.1016/

j.palaeo.2005.02.013.

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